A New Species of the Genus Gobio Cuvier, 1816 (Actynopterigii, Cyprinidae) from the Iberian Peninsula and Southwestern France

Graellsia, 60(1): 107-116 (2004)

A NEW SPECIES OF THE GENUS GOBIO CUVIER, 1816 (ACTYNOPTERIGII, CYPRINIDAE) FROM THE IBERIAN PENINSULA AND SOUTHWESTERN FRANCE

I. Doadrio1 and M. J. Madeira2

ABSTRACT

Iberian and southern French populations of the genus Gobio, considered in the past to be populations of Gobio gobio, are assigned to a new species (Gobio lozanoi n. sp.) based on genetic and morphological characters. This new species of the genus Gobio is found in the basins of the Rivers Adour in France and Bidasoa, Duero, Ebro, Guadalete, Guadiana, Guadalquivir, Júcar, Llobregat, Mondego, Mijares, Nalón, Nansa, Miño, Segura, Tajo, and Turia in the Iberian Peninsula. The new species is distinguished from Gobio gobio by a combination of the following characters: 36-39 scales on the lateral line, 3 scales below the lateral line. The distance between the pectoral and ventral fins is greater than that between the ventral and anal fins (VAL/PVL≤0.8). The preorbital distance is short and the head is wide (ED/PrOL≥0.56; PrOL/HH≤0.69; PrOL/HW≤0.68). Divergence distances in cytochrome b between Gobio gobio and the new species are “p”=4.8-5.9%. Key words: Cypriniformes, Cyprinidae, Taxonomy, Gobio gobio, Iberian Peninsula, South France.

RESUMEN Una nueva especie del género Gobio Cuvier, 1816 (Actinopterygii, Cyprinidae) de la Península Ibérica y sur de Francia Se describe una nueva especie del género Gobio de la Península Ibérica y sur de Francia (Gobio lozanoi n. sp.), considerada hasta ahora una población de Gobio gobio, en base a caracteres genéticos y morfológicos. La nueva especie del género Gobio pro- cede de las cuencas de los ríos Adour en Francia y Bidasoa, Duero, Ebro, Guadalete, Guadiana, Guadalquivir, Júcar, Llobregat, Mondego, Mijares, Nalón, Nansa, Miño, Segura, Tajo y Turia en la Península Ibérica. Esta nueva especie se diferencia de Gobio gobio por una combinación de los siguientes caracteres: 36-39 escamas en la línea lateral, 3 escamas por debajo de la línea lateral. La distancia entre la aleta pectoral y ventral es siempre mayor que entre la aleta ventral y la anal (VAL/PVL≤0.8). La distancia preorbital es corta y la cabeza ancha (ED/PrOL≥0.56; PrOL/HH≤0.69; PrOL/HW≤0.68). Para el citocromo b la divergencia entre Gobio gobio y la nueva especie fue de “p”=4.8-5.9%. Palabras clave: Cypriniformes, Cyprinidae, Taxonomía, Gobio gobio, Península Ibérica, Sur de Francia.

1 Museo Nacional de Ciencias Naturales. José Gutiérrez Abascal 2. 28006 Madrid. Spain. [email protected] 2 Dep. Zoología y Dinámica Celular Animal. Fac. Farmacia. Universidad del País Vasco (U.P.V). Paseo de la Universidad 7. 01006 (Vitoria-Gasteiz) Spain. [email protected]

Introduction ses previously, it had not been named until this study was made. Gobio (Cuvier, 1816) is a genus of approxima- tely 20 species distributed throughout Europe and Asia. Within this genus, Gobio gobio (Linnaeus, Material and Methods 1758) with its numerous described subspecies and local forms, is one of the most variable fish species The description of this new Gobio species is in all of Europe owing to its phenotypic plasticity. based on the study of sixty-three adults from the The common gudgeon (Gobio gobio) is a Eresma River in Coca, Segovia. The holotype and a Palaearctic species whose distribution area series of paratypes were deposited in Museo extends throughout most of Europe, Anatolia, Nacional de Ciencias Naturales (Spain). For the Siberia and Central Asia. In the Iberian Peninsula morphometric analysis we studied the following it was cited for the first time in Matosinho, Gobio gobio material: MNCN20568-20573, six Portugal (Osorio, 1896) and later in the Duero individuals from Vltava River, Brno, Czech basin in Spain (Lozano-Rey, 1919). Its dispersal Republic. MNCN20698-20707, 10 individuals capacity and ability to colonize new areas favou- from Elbe River, Cernylev, Czech Republic. red its rapid expansion throughout the entire geo- MNCN20588-20594, 7 individuals from Elbe graphical area of the Iberian Peninsula over the River, Nova Ves, Czech Republic. 94253-94263, 11 last decades. As a result, the common gudgeon individuals from Mures River Lunca-Bradului, was considered to be an introduced species in the Romania. MNCN94322-94335, 14 individuals Iberian Peninsula. Only one study, based on bio- from Mures River Brancovenesti (Romania). metrical analyses, considered the allochthonous MNCN195547-195556, 10 individuals from Tha- nature of gudgeon populations in the Iberian mes River, Lombard Road, England. Peninsula (Coelho, 1981). Twenty-five morphometric variables were mea- However, in the course of recent genetic studies sured. All measurements were in millimetres and of different species of the genus Gobio, using a were log-transformed for morphometric analysis. mitochondrial DNA (cytochrome b) marker, the G. The following abbreviations were used for gobio populations from the Iberian Peninsula and morphometric and meristic characters: SL, stan- southern France (Adour basin) were found to be dard length; HL, head length; HH, head height; genetically different from G. gobio populations in HW, head width; PrOL, preorbital length; ED, eye England and Central Europe (Madeira et al., in diameter; ID, interorbital distance; PRDD, predor- press). This last study confirms a phylogeographic sal distance; PrPD, prepectoral distance, PrVD, genetic discontinuity between Iberian and other preventral distance; PrAD, preanal distance; CPL, European populations. Thus, Iberian and Adour caudal peduncle length; APL, anal peduncle Basin G. gobio populations seem to have endemic length; PVL, pectoral-ventral length; VAL, ven- origins. However, the majority of the endemic tral-anal length; DFL, dorsal fin length; DFH, dor- freshwater fishes of the Iberian Peninsula have sal fin height; PFL, pectoral fin length; VFL, narrower distributions ranges. ventral fin length ; AFL, anal fin length; AFH, anal Gobio gobio populations from the Iberian fin height; CFL, caudal fin length, BD body depth; Peninsula have a decidedly wider distribution BLD, body least depth; BL, barbel length; D, dor- range, and have been reported in all the main river sal fin rays; A, anal fin rays; P, fin rays; V, fin rays; basins of Spain (Doadrio & Elvira, 1986) and C, fin rays; LLS, lateral line scales; LTU, upper Portugal (Almaça, 1965). This may be due to its use transversal scales; LTL, lower transversal scales; as live-bait in sport fishing and to having been PT, pharyngeal teeth. Only branched fins rays were translocated from basin to basin by human activity. counted. As a consequence, there is low genetic variation Differences in body shape between populations between the populations from the Iberian Peninsula were analysed using Principal Component Analysis and southern France (Adour basin), making it diffi- (PCA). The Shear method was used to correct size cult to determine in what basin the native popula- effects (SPCA) (Humphries et al., 1981) using the tions originated. programme SHEAR PCA available from N. The purpose of this study was describe the spe- MacLeod Natural History Museum London cimens from the Iberian Peninsula and Adour Basin (http//www.nhm.ac.uk/hosted_sites/paleonet/ftp/ftp.html). as new species of the genus Gobio. Although this Institutional acronyms: MNCN Museo Nacional species had been defined through molecular analy- de Ciencias Naturales.

Fig. 1.— Gobio lozanoi sp. nov. Holotype MNCN153558; Eresma River, Duero Basin, Coca, Segovia. Spain. Scale = 10 mm.

Fig. 1.— Gobio lozanoi sp. nov. Holotype MNCN153558; río Eresma, Cuenca del río Duero, Coca, Segovia. España. Escala = 10 mm.

Gobio lozanoi, new species 1.4-1.8 (1.5) times the head height and 1.45-1.82 (Fig. 1) (1.61) the width head (Table 3a-c). The eye diameter length is 1.4-2.1 (1.8) times the preorbital HOLOTYPE. MNCN153558, 88.2 mm SL; Eresma River, Duero Basin, Coca, Segovia. Spain. Leg J. Lobón, S. Torres and I. distance. Ventral fin is inserted posteriorly to the Doadrio. 14.VIII.1984 (Table 1). origin of the dorsal fin, on the same axis (Table 3d). Predorsal length is 1-1.1 (1.1) times the pre- PARATYPES. MNCN153557, 153559-153620. Eresma River, ventral length. Distance between pectoral and Duero Basin, Coca, Segovia. Spain. Leg J. Lobón, S. Torres and I. Doadrio. 14.VIII.1984. ventral fin is longer than ventral-anal distance. Pectoral-ventral distance is 1.1-1.5 (1.2) times DIAGNOSIS. Differs from all other known species ventral-anal distance. Fin size moderately large. of Gobio by the following: 36-39 scales in the late- There is a pair of maxillary barbels of variable ral line, 3 scales below the lateral line, distance bet- size. All populations of this species except those ween pectoral and ventral fin is greater than ventral from the Bidasoa River in the Iberian Peninsula anal distance (VAL/PVL≤0.8), the preorbital dis- and those from the Adour River in France have tance is short and the head is wide (ED/PrOL≥0.56, longer barbels than G. gobio. Barbels reach the PrOL/HH≤0.69; PrOL/HW≤0.68). Divergence dis- posterior border of the eye. Scales larger than tances in cytochrome b between G. gobio and the those of G. gobio and there are only 3 scales on new species are “p”=0.48-0.59. the transversal rows under the lateral line. DESCRIPTION. D II-III 7, A II-III 6, P 14 (15), V PIGMENTATION PATTERN. One row of 6-12 (7) II 6 (7), C, LLS 36-39 (x =37.4), LTU 5-6 (x = dark blotches along the lateral line. Except in the 5.3), LTL 3, PT 5.3 (2)-5.3 (2), 18 (17) abdominal ventral part of the body the scales are black remar- vertebrae, 17 caudal vertebrae. Morphometric ked. Upper cephalic region darkly spotted. Below characters are given in Tables 1 and 2. The body is the eye there is a characteristic elongated black elongated and moderately compressed. Maximum spot. The dorsal, caudal, pectoral and anal fins have body depth is 4-5.2 (4.5) times the standard a variable number of rows of dark spots. Above the length. Minimum body depth is 3.7-5.2 (4.4) times lateral line the body is grey or brown while below the length of the dorsal caudal peduncle. The head it is light brown. is wider and shorter than that of G. gobio and is ETYMOLOGY. The species is dedicated to Luis 3.8-4.3 (4) times the standard length. Preorbital Lozano Rey (1878-1958) for his contribution to our distance short, 2.3-3 (2.5) times the head length; knowledge of Iberian freshwater fishes.

Table 1.— Statistical parameters for the morphometric and Table 2.— Statistical parameters for the morphometric charac- meristic characters of G. lozanoi sp. nov. type series. Variables ters of G. lozanoi sp. nov. Each variable is divided by standard are described in Methods. (SD = Standard deviation). length. Variables are described in Methods. (SD = Standard deviation). Tabla 1.— Parámetros estadísticos para los caracteres morfo- métricos y merísticos de la serie tipo de G. lozanoi sp. nov. Las Tabla 2.— Parámetros estadísticos para los caracteres morfo- variables se describen en la metodología. (SD = desviación métricos de G. lozanoi sp. nov. Cada variable está dividida por estandard). la longitud estandar. Las variables se describen en la metodo- logía. (SD = desviación estandard).

Gobio lozanoi nov. sp. Gobio lozanoi nov. sp. (n= 63) Variable Holotype Paratypes (n= 62) Range Mean SD Variable Mean Range SD SL 88.2 57.1-114.6 88.1 10.6 SL 88.07 57.10-114.6 11.41 HL 22.8 15-28.3 22.1 2.5 HL/SL 0.25 0.23-0.26 0.01 PrOL 9.2 4.9-12.2 8.9 1.3 PrOL/SL 0.10 0.09-0.11 0.01 ED 4.9 3.3-6.2 5 0.6 ED/SL 0.06 0.05-0.07 0.00 ID 6.2 4.2-10.2 6.5 1.1 ID/SL 0.07 0.06-0.09 0.01 HW 14.3 8-18 14.2 1.9 HW/SL 0.16 0.14-0.18 0.01 HH 15.5 8.5-17.7 13.6 1.8 HH/SL 0.15 0.14-0.18 0.01 PrPD 21.9 14.8-28.1 22 2.7 PrPD/SL 0.25 0.23-0.27 0.01 PrVD 44.8 30.2-56.2 44.3 5.1 PrVD/SL 0.50 0.47-0.53 0.01 PRAD 60.9 39.6-77.3 61.8 7.5 PRAD/SL 0.70 0.66-0.73 0.01 PrDD 41.0 26.4-54.2 41.8 5 PrDD/SL 0.47 0.45-0.5 0.01 CPL 39.3 23.1-46.6 37.4 4.8 CPL/SL 0.42 0.39-0.46 0.02 APL 20.1 13.6-25.8 19.7 2.3 APL/SL 0.22 0.19-0.24 0.01 PVL 19.6 13.-26.5 19.9 2.4 PVL/SL 0.23 0.21-0.25 0.01 VAL 15.9 9.5-22.7 16 2.4 VAL/SL 0.18 0.16-0.21 0.01 DFL 11.3 7.5-14.8 11.5 1.6 DFL/SL 0.13 0.11-0.15 0.01 DFH 19.3 13.9-25 18.9 2.1 DFH/SL 0.22 0.19-0.24 0.01 PFL 19.8 13-23 18.9 2.2 PFL/SL 0.22 0.19-0.25 0.01 VFL 15.8 10.5-19.5 15.7 1.7 VFL/SL 0.18 0.16-0.2 0.01 AFL 6.6 4.3-8.5 6.6 0.9 AFL/SL 0.07 0.06-0.09 0.01 AFH 14.7 10-19.4 14.5 1.9 AFH/SL 0.16 0.14-0.19 0.01 CFL 15.7 11.8-20.2 16.6 1.8 CFL/SL 0.19 0.16-0.21 0.01 BD 21.2 12.2-26 19.7 2.6 BD/SL 0.22 0.19-0.25 0.01 BLD 8.5 5.4-11.3 8.5 1.2 BLD/SL 0.10 0.08-0.11 0.01 BL 8.9 4.6-11.1 8.6 1.2 BL/SL 0.10 0.08-0.12 0.01 LLS 37 36-39 37.4 0.8 LTU 5 5-6 5.3 0.4 LTL 3 3-3 3 0 D 7 7.7 7 0 genus Gobio to be allochthonous (Lozano-Rey, A 6 6-6 6 0 1935, Coelho, 1981, Lobón-Cerviá & Torres, 1984, Doadrio & Elvira 1986, Lobón-Cerviá et al., 1991). The absence of reliable mentions prior to 1913 for Iberian G. gobio (Lozano Rey, 1919), a fact that is DISTRIBUTION. Currently G. lozanoi is present in truly unusual considering that today it is one of the the Iberian Peninsula and in southern France in the most abundant species in the Iberian Peninsula, basins of the Rivers Adour, Bidasoa, Duero, gives credence to an allocthonous origin. Mondego, Ebro, Guadalete, Guadiana, Guadalquivir, The first reference to the genus Gobio in the Júcar, Llobregat, Mijares, Nalón, Nansa, Miño, Iberian Peninsula is from Matosinho, a village loca- Segura, Tajo and Turia (Fig. 2). For many years, aut- ted on the Atlantic coast near the estuary of the hors have considered Iberian populations of the Duero River in Portugal (Osorio, 1896). Today, the

Fig. 2.— Distribution range of G. lozanoi sp. nov.

Fig. 2.— Área de distribución de G. lozanoi sp. nov.

Table 3.— Frequency distribution of preorbital indexes: A) PrOL/ED; B) PrOL/HH; C) PrOL/HW and ventral index: D) VAL/PVL in G. gobio and G. lozanoi sp. nov.

Tabla 3.— Distribución de frecuencias de los índices preorbitales: A) PrOL/ED; B) PrOL/HH; C) PrOL/HW e índice ventral: D) VAL/PVL en G. gobio y G. lozanoi sp. nov.

A) PrOL/ED Species 0.3-0.40 0.41-0.50 0.51-0.60 0.61-0.70 0.70-0.80 Gobio lozanoi n. sp. 054882 Gobio gobio 26 32

B) PrOL/HH Species 0.51-0.60 0.61-0.70 0.71-0.80 0.81-0.90 0.91-1 Gobio lozanoi n. sp. 75600 Gobio gobio 0 0 33 22 3

C) PrOL/HW Species 0.55-0.60 0.61-0.65 0.66-0.70 0.71-0.75 0.76-0.80 0.81-0.85 0.86-0.9 Gobio lozanoi n. sp. 18 29 16 0 Gobio gobio 002251663

D) VAL/PVL Species 0.70-0.75 0.76-0.8 0.81-0.85 0.86-0.90 0.91-0.95 0.96-1 1.05-1.1 1.11-1.15 Gobio lozanoi n. sp. 12 17 22 12 0 0 0 Gobio gobio 0 0 0 0 13 17 13 2

.179

.144 + #

| | G # .110 + | G # | ### | GGG ## | GGG G G R | GGGG#### .076 + GGRG R ### | G GG GGG G ### GGR GGG #R# ## PC 2 | | RRR GG ## # # | G RGG ###R # | GGRG ## ## .041 + GG G ## | G ## # # | # | G | G | # .007 + | #

-.028 .150 .231 .313 .394 .191 .272 .353

PC 3

Fig. 3.— Bivariate Plot of Principal Components 2 VS. Principal Components 3 for 24 Shear corrected variables. l Gobio lozanoi (n= 63). H Gobio gobio (n= 58). R = Multiple Individual Location. + = Group Centroid

Fig. 3.- Gráfico bivariante del segundo componente principal frente al tercero para 24 variables corregidas mediante Shear. l Gobio lozanoi (n= 63). H Gobio gobio (n= 58). R = Localizaciones individuales múltiples + = Centroide. species is not found in Matosinho nor is there an Iberian populations located in the Bidasoa and appropriate habitat for the genus Gobio in this Ebro Rivers. Recently, molecular analysis has region. It is likely, therefore, that Osorio’s (1896) revealed that Iberian and French populations original citation refers to the genus Gobius; a genus from various drainages are autochthonous and that is often found in estuaries and marine environ- originated from a limited area of the Iberian ments. The first reliable locality reference for the Peninsula or south of France (Madeira et al., in genus Gobio is the Voltoya River, in the Duero basin, press). from which individuals were deposited in the Further genetic variability studies carried out in MNCN in 1913 (Lozano-Rey, 1919, Doadrio, 1989). the different basins could resolve the question of Just within the last years Doadrio (2001) has where exactly in the Iberian Peninsula and southern proposed an autochthonous origin for some of the France the species originated.

Table 4.— Eigenvectors for the first three principal compo- COMMON NAMES. Gobio, Cabezudo, Gobi, nents shear transformed for 24 variables. Gobioa. Tabla 4.— Eigenvectors para los tres primeros componentes REMARKS. The species inhabits very different principales transformados por el método de Shear. kind of habitats. In northern Spain, southern France and the Duero basin it characteristically inhabits streams and rivers with clear water, gravel or sandy Eigenvectors I II III bottoms. In the central Iberian Peninsula this spe- SL 0.109849 0.059693 0.022847 cies also lives in lowland waters that are characte- HL 0.148667 0.030301 0.025726 ristically slow moving with gravel, sandy and PrOL 0.150266 0.125809 0.032930 clayey bottoms. The spawning period is longer in G. lozanoi populations than in G. gobio (Lobón- ED 0.222007 0.374092 0.052206 Cerviá & Torres, 1984). In the Lozoya River, spaw- HW 0.393193 -0.792203 -0.037895 ning occurrs from mid May and early June to the HH 0.230504 -0.014518 0.131641 end of July or early August (Lobón-Cerviá & PrPD 0.214581 -0.043235 -0.104643 Torres, 1984) and from June to August in the PrVD 0.158466 0.122240 0.020712 Nivelle River (Bernet, 1960). The life span of G. PRAD 0.132064 0.050397 0.069842 lozanoi is short; reproduction is early (1-year-old) PrDD 0.125980 0.056656 0.052370 and tends to be semelparous (Lobón-Cerviá et al., CPL 0.133494 0.067586 0.002887 1991). Populations of G. lozanoi are highly abun- APL 0.133935 0.093201 -0.009521 dant, and in the Ucero River (Duero basin) densi- PVL 0.132788 0.197345 0.056381 ties of 1047 individuals per hectare have been VAL 0.186124 -0.119967 0.116260 reported (Lobón-Cerviá et al., 1986) DFL 0.167926 0.215331 0.123474 DFH 0.225445 0.067765 -0.230068 PFL 0.145541 0.026126 -0.013758 Discussion VFL 0.151991 0.004140 -0.058898 AFL 0.168052 -0.071337 -0.034807 Populations from the Iberian Peninsula and AFH 0.296228 0.147692 -0.744656 Adour River in France are so highly divergent from CFL 0.177266 0.034647 -0.033995 other populations of G. gobio, both morphologi- BD 0.203126 -0.052513 -0.037592 cally and genetically (Table 5), that separating them BLD 0.298598 -0.144439 0.206496 into distinct species is warranted. No existing name ID 0.304277 0.120158 0.517391 can be assigned to them; so therefore, we have cho- sen to call this population Gobio lozanoi sp. nov. Gobio lozanoi is differentiated from other populations of G. gobio by: 1) its much shorter snout

Table 5.— Summary of diagnostic characters of G. lozanoi sp. nov.

Tabla 5.— Resumen de caracteres diagnósticos de G. lozanoi sp. nov.

Characters G. gobio (n= 58) G. lozanoi sp. nov. (n= 63)

Number of scales in the lateral line x =41 (39-43) x = 37.4 (36-39) Number of scales below of the lateral line 4 3 Preorbital index (PrOL/HH) 0.5-0.7 0.7-1 Ventral Index (VAL/PVL) 0.7-0.9 0.9-1.1 Divergence to cytochrome b gene sequences “p”=4.8-5.9%

Fig . 4 R. rutilus P. parva B. haasii G. ciscaucasicus 100 G. uranoscopus G. gobio (Danube ) 52 G. lozanoi (Tajo) G. lozanoi (Ebro) G. lozanoi (Ebro) G. lozanoi (Duero ) 100 G. lozanoi (Duero) G. lozanoi (Ebro) G. lozanoi (Ebro) G. lozanoi (Ebro) G. lozanoi (Tajo) G. lozanoi (Ebro) 100 G. lozanoi (Ebro) G. lozanoi ( Duero) G. lozanoi (Tajo) G. lozanoi (Ebro) G. lozanoi (Ebro) G. lozanoi ( Bidasoa) G. lozanoi ( Bidasoa) G. lozanoi ( Ebro) G. lozanoi (Ebro) G. lozanoi ( Duero) 99 G. lozanoi ( Duero) G. lozanoi (Adour) G. lozanoi ( Adour) G. lozanoi ( Adour) G. lozanoi (Ebro) G. lozanoi ( Ebro) G. lozanoi (Adour) G. lozanoi ( Duero) G. lozanoi (Nansa) G. lozanoi (Adour) G. lozanoi (Adour) 0.005 substitutions/site

Fig. 4.— Phylogenetic tree of 35 analysed specimens of the genus Gobio recovered from cytochrome b sequences according to the neighbour-joinig method based on the best model of evolution that fit our data using the program Model test 3.04 (Posada & Crandall, 1998). Branch lengths are proportional to the estimated mean number of substitutions per site (see scale bar) (from Madeira et al., in press). The Ibero-French populations are here described as G. lozanoi n. sp. Numbers are bootstrap values for 500 replicates.

Fig. 4.— Árbol filogenético para 35 individuos del género Gobio basado en secuencias de citocromo b y realizadas con el méto- do de “neighbour-joinig” basado en el mejor modelo evolutivo que se ajustó a nuestros datos usando el programa Model test 3.04 (Posada & Crandall, 1998). La longitud de las ramas es proporcional al número medio de sustituciones por sitio (ver escala) (toma- do de Madeira et al., in press). Las poblaciones ibero-francesas son descritas aqui como G. lozanoi n. sp. Números son valores de Bootstrap para 500 réplicas.

(c) Sociedad de Amigos del Museo http://graellsia.revistas.csic.es Nacional de Ciencias Naturales y Consejo Superior de Investigaciones Científicas Licencia Creative Commons 3.0 España (by-nc) A NEW SPECIES OF THE GENUS GOBIO 115 and wide head 2) fewer scales on and below the Beall, C. García de Leaniz, J. Álvarez and the Guard Services lateral line 3) greater length distance between pec- of the Navarra and Gipuzkoa local governments. Last, we are grateful to A. Machordom, B. Gómez-Moliner and R. Zardoya toral and ventral fins than between ventral-anal. for their suggestions and improving some of the English text. A PCA analysis with 24 Shear morphometric James Watkins further revised the English text. corrected variables was carried out to remove size effect. The barbels length was removed because of the high variability found in this character in the References Iberian populations. The area of the scores of each component differed significantly (MANOVA) bet- ALMAÇA, C., 1965. Contribution à la connaissance des ween G. gobio and G. lozanoi for the first three poissons des eaux intérieures du Portugal. Revista da components. Significant differences in morphology Facultade Ciências. Lisboa. 2ª serie, 13(2): 225-262. were found between G. lozanoi and G. gobio prin- BANARESCU, P., SORIC, M. V. & ECONOMIDIS, P. S., 1999. cipally as a result of the contribution of the second Gobio gobio (Linnaeus, 1758). In: P. Banarescu size corrected component (Fig. 3). The highest (ed.). The Freshwater fishes of Europe, vol. 5(I). eigenvector for the second component was the wide Aula Verlag. Wiesbaden: 78-134. head (Table 4). BERNET, B., 1960. Recherches biologiques sur les popu- The Gobio gobio with the least number of sca- lations de Gobio gobio (Linné, 1758) de la Nivelle, les on the lateral line (x =37.5 36-39) (Banarescu fleuve cotier du Pays Basque. Annales de la Station et al., 1999) are found in populations from Skadar Centrale d’Hydrobiologie Appliquée, 8: 127-180. lake (x =38.5 33-41; Ivanovic, 1973) and the Po BIANCO, P. G. & TARABORELLI, T., 1984. Gobio gobio River (x =38.4 37-40; Bianco & Tabborelli, 1984 benacensis (Pollini, 1816) sottoespecie valida per Notwithstanding, we found that G. gobio always l’Italia (Pisces, Cyprinidae). Bolletino Museo Civico has a greater number of scales on the lateral line di Storia Naturale Verona, 11: 525-536. than does G. lozanoi. Three scales on the lower row COELHO, M. M., 1981. Contributions to the knowledge of the lateral line have been found before in Iberian of the populations of Gobio gobio (Linnaeus, 1758) populations (Coelho, 1981) but not in other (Pisces, Cyprinidae) in Portugal. Arquivos do Museo European populations (Banarescu et al., 1999). Bocage, 1(5): 67-94. Head and preorbital lengths reported in the literatu- DOADRIO, I., 1989. Catálogo de los peces de agua dulce re for G. gobio populations are always greater than del Museo Nacional de Ciencias Naturales. MNCN- those reported for G. lozanoi. Only some popula- CSIC. Madrid. 185 pp. tions of G. gobio lepidolaemus and G. gobio kovat- DOADRIO, I., 2001. Atlas y libro rojo de la ictiofauna con- chevi have similar values to G. lozanoi (Banarescu tinental española. MIMAM-CSIC. Madrid. 363 pp. et al., 1999). DOADRIO, I. & CARMONA, J. A. (In press). Phylogenetic Within of genus Gobio the sister species of relationships of the genus Chondrostoma Gobio lozanoi is G. gobio (fig. 4) other european (Actynopterygii, Cyprinidae) using cytochrome b species of this genus are very different morphologi- sequences Molecular Phylogenetics and Evolution. cally and belong to the subgenera Romanogobio or DOADRIO, I. CARMONA, J. A. & MACHORDOM, A., 2002. Rheogobio (Banarescu et al., 1999). The genetic Haplotype diversity and phylogenetic relationships among the Iberian Barbels (Barbus, Cyprinidae) differentiation between G. gobio and G. lozanoi reveal two evolutionary lineages. The Journal of was “p”=4.8-5.9%; (Madeira et al. in press) a Heredity, 93(2): 140-147. divergence similar to that found between other DOADRIO, I. & ELVIRA, B., 1986. Sobre la distribución de Iberian species of the genera Barbus (Doadrio et Gobio gobio (L., 1758) (Ostariophysi, Cyprinidae) al., 2002), Chondrostoma (Doadrio and Carmona, en España. Doñana Acta Vertebrata, 13: 165-166. in press) and Squalius (Sanjur et al., 2003). HUMPHRIES, J. M., BOOKSTEIN, F. L., CHERNOFF, C., SMITH, G. R., ELDER, R. L. & POSS, S. G., 1981. Multivariate discrimination by shape in relation to size. Systematic AKNOWLEDGEMENTS Zoology, 30: 291-308. This study was financed by the DGES project REM2001- IVANOVIC, B. M., 1973. Ichthyofauna of Skadar Lake. 0662/G10. We thank J. Cubo, I. P. Garzón, J. A. González- Biological Station. Titograd. 146 pp. Carmona and A. Perdices for their assistance with field LOBÓN-CERVIÁ, J., DE SOSTOA, A. & MONTAÑÉS, C., collections. We are grateful to the Basque Country Goverment who supported the initial project about the species Gobio gobio 1986. Fish production and its relation with the com- in the Bidasoa basin and to the people who helped us with fish munity structure in an aquifer-fed stream of old collection for the initial study: R. Asensio, I. Urrizalki, E. Castile (Spain). Polskie Archiwum für Hydrobiologii, 33(3/4): 333-343.

LOBÓN-CERVIÁ, J., MONTAÑÉS, C. & DE SOSTOA, A., POSADA, D. & CRANDALL, K. A., 1998. Modeltest: tes- 1991. Influence of environment upon the life history ting the model of DNA substitution. Bioinformatics, of gudgeon Gobio gobio (L.): a recent and successful 14: 817-818 colonizer of the Iberian Peninsula. Journal of Fish OSORIO, B., 1896. Peixes de Matosinhos (Terceiro Biology, 39: 285-300. appendice ao catalogo dos peixes de portugal de LOBÓN-CERVIÁ, J. & TORRES, S., 1984. On the growth and Felix Capello. Jornal de Sciencias Mathematicas, reproduction of two populations of gudgeon (Gobio Physicas e Naturaes 2ª serie, 15: 131-159. gobio L.) in Central Spain. Acta Hydrobiologica, SANJUR, O. I., CARMONA, J. A. & DOADRIO, I. 2003. 25/26(1): 101-115. Evolutionary and biogeographical patterns within LOZANO-REY, L., 1919. Los peces de la fauna ibérica en Iberian populations of the genus Squalius inferred la colección del Museo el 1 de enero de 1919. from molecular data. Molecular Phylogenetics and Trabajos del Museo Nacional de Ciencias Naturales. Evolution, 29: 20-30. Madrid (serie Zool). 39: 1-112. LOZANO-REY, L., 1935. Los peces fluviales de España. Memorias de la Real Academia de Ciencias Exactas, Físicas y Naturales, 5: 1-390 MADEIRA, M. J., GÓMEZ-MOLINER, B. J. & DOADRIO, I. (In press). Genetic characterization of Gobio gobio Recibido, 15-V-2004 populations of the Iberian Peninsula based on Aceptado, 5-VII-2004 cytochrome b sequences. Folia Zoologica. Publicado, 25-VIII-2004

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